La. Mcdade et Md. Turner, STRUCTURE AND DEVELOPMENT OF BRACTEAL NECTARY GLANDS IN APHELANDRA (ACANTHACEAE), American journal of botany, 84(1), 1997, pp. 1-15
A survey of bracteal (extrafloral) nectaries in species of Aphelandra
(Acanthaceae) reveals substantial diversity. Each bracteal nectary is
an aggregate of individual glands that vary in number, size, and struc
ture among species. Glands contain three cell layers: a palisade-like
secretory cell layer, a one-to-many-celled intermediate layer with thi
ckened cell walls, and a foot layer Members of the A. pulcherrima comp
lex have one of two distinct gland types: relatively small glands with
a single-celled intermediate layer or larger glands that have a multi
cellular intermediate layer. Nectaries composed of small glands are pa
tches of many (>50) glands, whereas those composed of large glands are
patches of <10 glands. Four outgroup species have bracteal nectaries
of numerous small glands with pluricellular intermediate layers. Gland
s of all three types are initiated as single enlarged protodermal cell
s, and all undergo similar early periclinal divisions; the large-gland
type shows greater subsequent enlargement with many more anticlinal d
ivisions. The bracteal nectar glands are interpreted to be homologous
with simpler glandular trichomes, and mark a monophyletic lineage with
in Aphelandra. Comparisons with outgroup species show that both nectar
y types in the A. pulcherrima complex have diverged from an ancestral
condition of numerous small glands with pluricellular intermediate lay
ers. Use of the ontogenetic criterion to polarize gland type within th
e A. pulcherrima complex would yield erroneous results because evoluti
on has apparently involved a developmental truncation with loss of cel
l divisions in the intermediate layer of small glands. Comparable nect
ar glands in more distant taxa are interpreted as remarkable cases of
convergent evolution, perhaps from similar trichome precursors.